''Horneophyton'' is an extinct early plant which may form a "missing link" between the

hornwort Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...

s and the

Rhyniopsida The rhyniophytes are a group of extinct early vascular plants that are considered to be similar to the genus ''Rhynia'', found in the Early Devonian (around ). Sources vary in the name and rank used for this group, some treating it as the class ...

. It is a member of the class

Horneophytopsida The Horneophytopsida, informally called horneophytes, are a class of extinct plants which consisted of branched stems without leaves, true roots or vascular tissue, found from the Late Silurian to the Early Devonian (around ). They are the sim ...

. ''Horneophyton'' is among the most abundant fossil organisms found in the

Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...

, a

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...

Lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...

Aberdeenshire Aberdeenshire ( sco, Aiberdeenshire; gd, Siorrachd Obar Dheathain) is one of the 32 council areas of Scotland. It takes its name from the County of Aberdeen which has substantially different boundaries. The Aberdeenshire Council area inclu ...

, UK. A single species, ''Horneophyton lignieri'', is known. Its probable female

gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...

is the form taxon ''Langiophyton mackiei''.

Description

The

sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote ...

had bare stems (axes) up to 20 cm high and about 2 mm in diameter with an undivided cortex. Stomata were present but rare. There was a thin central strand of conducting tissue, but this was not reinforced with spiral and reticulate thickenings and thus does not constitute true

vascular tissue Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. The ...

. Early stages of development of the sporophytes of ''Horneophyton'' (as of hornworts) may have been dependent on their parent gametophytes for nutrition, but mature specimens have expanded, corm-like bases to their stems, up to 6 mm in diameter, that bore rhizoids and appear to be anchored in soil, suggesting a capacity for independent existence after the gametophyte had degenerated. The

sporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...

(spore-forming organ) is unique among both living and fossil plants, consisting as it does of branched lobes at the apex of some of the branches of the stem. Each lobe contains a central collumella, analogous to the sporangia of hornworts; however, the sporangia of hornworts are not branched. The number of lobes possessed by a sporangium varied; at least three orders of

dichotomous branching This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

have been found, resulting in more than four lobes. The sporangia were much less regular than shown in most reconstructions (including that opposite), and they had 'bumps' or emergences on them. Spores were released through a slit at the top of each lobe. The sporangia of ''Horneophyton'' contained trilete meiospores, the surfaces of which were decorated with short conical protuberances. The female

of the plant has been recognised and described as the form taxon ''Langiophyton mackiei''. It grew to a height of around 6 cm, and was free-living. The species was dioicous (unisexual), since it produced male and female

gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...

s on separate gametophytes. ''Horneophyton'' grew on sandy, organic-rich soil in damp to wet locations. They usually grew as isolated individuals. Rhynie chert with Horneophyton 1

Taxonomy

First named by Kidston & Lang in 1920 from

Early Devonian The Early Devonian is the first of three epochs comprising the Devonian period, corresponding to the Lower Devonian series. It lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, ...

fossils in the

, the original generic name ''Hornea'' was later found to occupied by a

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

in the family

Sapindaceae The Sapindaceae are a family of flowering plants in the order Sapindales known as the soapberry family. It contains 138 genera and 1858 accepted species. Examples include horse chestnut, maples, ackee and lychee. The Sapindaceae occur in temp ...

, ''Hornea mauritiana'', leading Barghoorn and Darrah to propose renaming the genus to ''Horneophyton'' in 1938. It was classified as a rhyniophyte (subdivision Rhyniophytina) by Banks, but the absence of true

led Kenrick and Crane in 1997 to create a new class,

, for this and similar genera. A single species, ''Horneophyton lignieri'', has been described.

Phylogeny

A possible phylogeny for ''Horneophyton'' is shown below (based on Crane et al. for the polysporangiophytes and Qiu et al. for the bryophytes. With vascular tissue but bryophyte-like vascular tissue and sporangia, the organism has been considered a missing link between the hornworts and the vascular plants or tracheophytes (which molecular data suggest are sister groups). Features suggesting a relationship with the hornworts include the general form of its sporangia; its corm also resembles the foot of some hornworts. The free living nature of its sporophytes, and the fact that they branching repeatedly, are marked differences which force it into the stem group of tracheophytes (along with ''Aglaophyton'').

References

External links

Horneophyton
at the University of Aberdeen. Includes images. (The apparent size of the corms in the reconstruction is inconsistent with the cross-sections shown earlier. See .)
Cladogram
from {{Taxonbar, from=Q148549 Early Devonian plants Prehistoric plant genera